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PARASITES

Parasitism is a type of symbiotic relationship between organisms of different species where one organism, the parasite, benefits at the expense of the other, the host. Traditionally parasite referred to organisms with lifestages that went beyond one host (e.g. Taenia solium), which are now called macroparasites (typically protozoa and helminths). Parasites can now also refer to microparasites, which are typically smaller, such as viruses and bacteria and can be directly transmitted between hosts of one species. Unlike predators, parasites are generally much smaller than their host, although both are special cases of consumer-resource interactions. Parasite show a high degree of specialization for their mode of life, and reproduce at a faster rate than their hosts. Classic examples of parasitism include interactions between vertebrate hosts and diverse animals such as tapeworms, flukes, the Plasmodium species, and fleas. Parasitism is differentiated from the parasitoid relationship, though not sharply, by the fact that parasitoids generally kill or sterilise their hosts. Parasitoidy occurs in about as many classes of organism as parasitism does. The harm and benefit in parasitic interactions concern the biological fitness of the organisms involved. Parasites reduce host fitness in many ways, ranging from general or specialized pathology (such as parasitic castration), impairment of secondary sex characteristics, to the modification of host behaviour. Parasites increase their fitness by exploiting hosts for resources necessary for the parasite's survival: (i.e. food, water, heat, habitat, and dispersal). Although the concept of parasitism applies unambiguously to many cases in nature, it is best considered part of a continuum of types of interactions between species, rather than an exclusive category. Particular interactions between species may satisfy some but not all parts of the definition. In many cases, it is difficult to demonstrate that the host is harmed. In others, there may be no apparent specialization on the part of the parasite, or the interaction between the organisms may be short-lived.

ETMOLOGY

First used in English 1539, the word parasite comes from the Medieval French parasite, from the Latin parasitus, the latinisation of the Greek (parasitos), "one who eats at the table of another"and that from (para), "beside, by" (sitos), "wheat".Coined in English in 1611, the word parasitism comes from the Greek (sitismos) "feeding, fattening".

TYPES

Parasites are classified based on their interactions with their hosts and on their life cycles. Parasites that live on the surface of the host are called ectoparasites (e.g. some mites). Those that live inside the host are called endoparasites (including all parasitic worms). Endoparasites can exist in one of two forms: intercellular parasites (inhabiting spaces in the host?s body) or intracellular parasites (inhabiting cells in the host?s body). Intracellular parasites, such as protozoa, bacteria or viruses, tend to rely on a third organism, which is generally known as the carrier or vector. The vector does the job of transmitting them to the host. An example of this interaction is the transmission of malaria, caused by a protozoan of the genus Plasmodium, to humans by the bite of an anopheline mosquito. Those parasites living in an intermediate position, being half-ectoparasites and half-endoparasites, are sometimes called mesoparasite. An epiparasite is one that feeds on another parasite. This relationship is also sometimes referred to as hyperparasitism, exemplified by a protozoan (the hyperparasite) living in the digestive tract of a flea living on a dog. Social parasites take advantage of interactions between members of social organisms such as ants or termites. In kleptoparasitism, parasites appropriate food gathered by the host. An example is the brood parasitism practiced by many species of cuckoo and cowbird, which do not build nests of their own but rather deposit their eggs in nests of other species and abandon them there. The host behaves as a "babysitter" as they raise the young as their own. If the host removes the cuckoo's eggs, some cuckoos will return and attack the nest to compel host birds to remain subject to this parasitism. The cowbird?s parasitism does not necessarily harm its host?s brood; however, the cuckoo may remove one or more host eggs to avoid detection, and furthermore the young cuckoo may heave the host?s eggs and nestlings out of the nest. Parasitism can take the form of isolated cheating or exploitation among more generalized mutualistic interactions. For example, broad classes of plants and fungi exchange carbon and nutrients in common mutualistic mycorrhizal relationships; however, some plant species known as myco-heterotrophs "cheat" by taking carbon from a fungus rather than donating it. Parasitoids are organisms whose larval development occurs inside or on the surface of another organism, resulting in the death of the host. This means that the interaction between the parasitoid and the host is fundamentally different from that of a true parasite and shares some of the characteristics of predation. An adelpho-parasite is a parasite in which the host species is closely related to the parasite, often being a member of the same family or genus. An example of this is the citrus blackfly parasitoid, Encarsia perplexa, unmated females of which may lay haploid eggs in the fully developed larvae of their own species. These result in the production of male offspring. The marine worm Bonellia viridis has a similar reproductive strategy, although the larvae are planktonic.

HOST DEFENSES

Hosts respond to parasitisms in many ways ranging from the morphological to the behavioural. In some cases, plants produce toxins to deter parasitic fungi and bacteria.Vertebrate animals have developed complex immune systems, which can target parasites through contact with bodily fluids. Animals are also known to resort to behavioral defenses, examples of which are the avoidance by sheep of open pastures during spring, when roundworm eggs accumulated over the previous year hatch en masse; and the ingestion of alcohol by infected fruit flies as self medication against blood-borne parasites. In humans, parasite immunity is developed prominently by Immunoglobulin E antibodies.

EVOLUTIONARY ASPECTS

Biotrophic parasitism is a common mode of life that has arisen independently many times in the course of evolution. Depending on the definition used, as many as half of all animals have at least one parasitic phase in their life cycles, and it is also frequent in plants and fungi. Moreover, almost all free-living animals are host to one or more parasites taxa. Restoration of a Tyrannosaurus with parasite infections. A 2009 study showed that holes in the skulls of several specimens might have been caused by Trichomonas-like parasites. Parasites evolve in response to the defense mechanisms of their hosts. As a result of host defenses, some parasites evolve adaptations that are specific to a particular host taxon, specializing to the point where they infect only a single species. Such narrow host specificity can be costly over evolutionary time, however, if the host species becomes extinct. Therefore many parasites can infect a variety of more or less closely related host species, with different success rates. Host defenses also evolve in response to attacks by parasites. Theoretically, parasites may have an advantage in this evolutionary arms race because of their more rapid generation time. Hosts reproduce less quickly than parasites, and therefore have fewer chances to adapt than their parasites do over a given span of time. In some cases, a parasite species may coevolve with its host taxa. Long-term coevolution sometimes leads to a relatively stable relationship tending to commensalism or mutualism, as, all else being equal, it is in the evolutionary interest of the parasite that its host thrives. A parasite may evolve to become less harmful for its host or a host may evolve to cope with the unavoidable presence of a parasite-- to the point that the parasite's absence causes the host harm. For example, although animals infected with parasitic worms are often clearly harmed, and therefore parasitized, such infections may also reduce the prevalence and effects of autoimmune disorders in animal hosts, including humans. Competition between parasites tends to favor faster reproducing and therefor more virulent parasites. Parasites whose life cycle involves the death of the host, to exit the present host and sometimes to enter the next, evolve to be more virulent or even alter the behavior or other properties of the host to make it more vulnerable to predators. Parasites that reproduce largely to the offspring of the previous host tend to become less virulent or mutualist, so that its hosts reproduce more effectively. The presumption of a shared evolutionary history between parasites and hosts can sometimes elucidate how host taxa are related. For instance, there has been dispute about whether flamingos are more closely related to the storks and their allies, or to ducks, geese and their relatives. The fact that flamingos share parasites with ducks and geese is evidence these groups may be more closely related to each other than either is to storks. Parasitism is part of one explanation for the evolution of secondary sex characteristics seen in breeding males throughout the animal world, such as the plumage of male peacocks and manes of male lions. According to this theory, female hosts select males for breeding based on such characteristics because they indicate resistance to parasites and other disease.

CO-SPECIATION

In rare cases, a parasite may even undergo co-speciation with its host. One particularly remarkable example of co-speciation exists between the simian foamy virus (SFV) and its primate hosts. In one study, the phylogenies of SFV polymerase and the mitochondrial cytochrome oxidase subunit II from African and Asian primates were compared. Surprisingly, the phylogenetic trees were very congruent in branching order and divergence times. Thus, the simian foamy viruses may have co-speciated with Old World primates for at least 30 million years.

ECOLOGY

QUANTITATIVE ECOLOGY

A single parasite species usually has an aggregated distribution across host individuals, which means that most hosts harbor few parasites, while a few hosts carry the vast majority of parasite individuals. This poses considerable problems for students of parasite ecology: the use of parametric statistics should be avoided. Log-transformation of data before the application of parametric test, or the use of non-parametric statistics is recommended by several authors. However, this can give rise to further problems. Therefore, modern day quantitative parasitology is based on more advanced biostatistical methods.

DIVERSITY ECOLOGY

Hosts represent discrete habitat patches that can be occupied by parasites. A hierarchical set of terminology has come into use to describe parasite assemblages at different host scales. Infrapopulation All the parasites of one species in a single individual host. Metapopulation All the parasites of one species in a host population. Infracommunity All the parasites of all species in a single individual host. Component community All the parasites of all species in a host population. Compound community All the parasites of all species in all host species in an ecosystem. The diversity ecology of parasites differs markedly from that of free-living organisms. For free-living organisms, diversity ecology features many strong conceptual frameworks including Robert MacArthur and E. O. Wilson's theory of island biogeography, Jared Diamond's assembly rules and, more recently, null models such as Stephen Hubbell's unified neutral theory of biodiversity and biogeography. Frameworks are not so well-developed for parasites and in many ways they do not fit the free-living models. For example, island biogeography is predicated on fixed spatial relationships between habitat patches ("sinks"), usually with reference to a mainland ("source"). Parasites inhabit hosts, which represent mobile habitat patches with dynamic spatial relationships. There is no true "mainland" other than the sum of hosts (host population), so parasite component communities in host populations are metacommunities. Nonetheless, different types of parasite assemblages have been recognized in host individuals and populations, and many of the patterns observed for free-living organisms are also pervasive among parasite assemblages. The most prominent of these is the interactive-isolationist continuum. This proposes that parasite assemblages occur along a cline from interactive communities, where niches are saturated and interspecific competition is high, to isolationist communities, where there are many vacant niches and interspecific interaction is not as important as stochastic factors in providing structure to the community. Whether this is so, or whether community patterns simply reflect the sum of underlying species distributions (no real "structure" to the community), has not yet been established.

ADAPTATION

Parasites infect hosts that exist within their same geographical area (sympatric) more effectively. This phenomenon supports the "Red Queen hypothesis?which states that interactions between species (such as host and parasites) lead to constant natural selection for adaptation and counter adaptation." The parasites track the locally common host phenotypes, therefore the parasites are less infective to allopatric (from different geographical region) hosts. Experiments published in 2000 discuss the analysis of two different snail populations from two different sources- Lake Ianthe and Lake Poerua in New Zealand. The populations were exposed to two pure parasites (digenetic trematode) taken from the same lakes. In the experiment, the snails were infected by their sympatric parasites, allopatric parasites and mixed sources of parasites. The results suggest that the parasites were more highly effective in infecting their sympatric snails than their allopatric snails. Though the allopatric snails were still infected by the parasites, the infectivity was much less when compared to the sympatric snails. Hence, the parasites were found to have adapted to infecting local populations of snails.

TRANSMISSION

Life cycle of Entamoeba histolytica, an anaerobic parasitic protozoan. Parasites inhabit living organisms and therefore face problems that free-living organisms do not. Hosts, the only habitats in which parasites can survive, actively try to avoid, repel, and destroy parasites. Parasites employ numerous strategies for getting from one host to another, a process sometimes referred to as parasite transmission or colonization. Some endoparasites infect their host by penetrating its external surface, while others must be ingested. Once inside the host, adult endoparasites need to shed offspring into the external environment to infect other hosts. Many adult endoparasites reside in the host?s gastrointestinal tract, where offspring can be shed along with host excreta. Adult stages of tapeworms, thorny-headed worms and most flukes use this method. Among protozoan endoparasites, such as the malarial parasites and trypanosomes, infective stages in the host's blood are transported to new hosts by biting-insects, or vectors. Larval stages of endoparasites often infect sites in the host other than the blood or gastrointestinal tract. In many such cases, larval endoparasites require their host to be consumed by the next host in the parasite's life cycle in order to survive and reproduce. Alternatively, larval endoparasites may shed free-living transmission stages that migrate through the host?s tissue into the external environment, where they actively search for or await ingestion by other hosts. The foregoing strategies are used, variously, by larval stages of tapeworms, thorny-headed worms, flukes and parasitic roundworms. Some ectoparasites, such as monogenean worms, rely on direct contact between hosts. Ectoparasitic arthropods may rely on host-host contact (e.g. many lice), shed eggs that survive off the host (e.g. fleas), or wait in the external environment for an encounter with a host (e.g. ticks). Some aquatic leeches locate hosts by sensing movement and only attach when certain temperature and chemical cues are present. Some parasites modify host behavior to make transmission to other hosts more likely. For example, in California salt marshes, the fluke Euhaplorchis californiensis reduces the ability of its killifish host to avoid predators. This parasite matures in egrets, which are more likely to feed on infected killifish than on uninfected fish. Another example is the protozoan Toxoplasma gondii, a parasite that matures in cats but can be carried by many other mammals. Uninfected rats avoid cat odors, but rats infected with T. gondii are drawn to this scent, which may increase transmission to feline hosts.

ROLES IN ECOSYSTEMS

Modifying the behavior of infected hosts, to make transmission to other hosts more likely to occur, is one way parasites can affect the structure of ecosystems. For example, in the case of Euhaplorchis californiensis (discussed above) it is plausible that the local predator and prey species might be different if this parasite were absent from the system. Although parasites are often omitted in depictions of food webs, they usually occupy the top position. Parasites can function like keystone species, reducing the dominance of superior competitors and allowing competing species to co-exist. Many parasites require multiple hosts of different species to complete their life cycles and rely on predator-prey or other stable ecological interactions to get from one host to another. In this sense, the parasites in an ecosystem reflect the "health" of that system.

EFFECTS ON HOST BEHAVIOR

Animal behavior is typically motivated by proximate level mechanisms that promote particular actions. In most cases, proximate level mechanisms are a result of the individual?s interaction with the environment but some parasites have been shown to interact and manipulate these mechanisms to produce behaviors beneficial to the parasite alone. Parasites are known to drastically affect how animals behave. Parasites most commonly target the central nervous system (CNS) in order to alter animal behavior. By affecting hormone secretions or by physical restructuring, parasites successfully change how an animal?s body functions and delivers, interprets and and reacts to messages. Some parasites, like toxoplasma, form vacuoles that travel through the nervous system interrupting key functions in intraneural communications. The emerald jewel wasp alters behavior through the injection of venom directly into the host?s brain, causing hypokinesia. Parasitic life cycles can have the capacity to infect a singular host or a series of hosts. Direct life cycles are related to a single host while indirect life cycles, or complex life cycles, rely on a series of intermediate hosts in order to complete the life cycle. Indirect life cycles likely arose in order to increase the efficiency of spreading to new hosts. By using intermediary?s the parasite DNA can spread to several hosts instead of remain concentrated within a single host, therefore promising a higher likelihood of DNA continuation. In some cases, intermediary hosts are accidental. Toxoplasma gondii is a noted case of unintended intermediary hosts. Typically, Toxoplasma infects animals from the Felidae family and oocysts are shed with the feces. When rodents consume the fecal matter, they become infected themselves and the parasite begins to alter their behavior. Rodents become more extroverted and less fearful of felines. Toxoplasma, however, has begun to infect humans. In doing so, human behavior has been altered in similar ways to rodents. Further, Toxoplasma has been linked to cases of schizophrenia. The emerald jewel wasp (Ampulex compressa) takes advantage of its host, the American cockroach (Periplaneta americana) specifically as a food source and home for its growing larvae. The wasp begins by injecting venom into the brain which paralyses the central nervous system system of the cockroach and puts it into a state of hypokinesia. Hypokinesia...is a reversible long-term lethargy characterized by lack of spontaneous movement or response to external stimuli, (Banks and Adams). After dragging the cockroach to a burrow, the wasp deposits an egg into its carcass, burying it for the growing larvae to feed off until it emerges in 6 weeks, leaving nothing but a hard outer cockroach shell. Although the circuitry in control of movement is functional, the nervous system acts from a depressed state. They are not killed by this hypokinesia, but would recover if not for the larvae eating them from the inside out. The movement is controlled by dopamine and octopamine which affect transmission of interneurons involved in the natural response to escape. Reduced motor activity results from a reduction of these amines.

VALUE

Although parasites are generally considered to be harmful, the eradication of all parasites would not necessarily be a noble aim. Among other things, parasites may account for as much as or more than half of life's diversity; they perform an important ecological role (by weakening prey) that ecosystems would take some time to adapt to; and without parasites organisms may eventually tend to asexual reproduction, dinimishing the diversity of sexually dimorphic traits. Parasites provide an opportunity for the transfer of genetic material between species. On rare but significant occasions this may provide a path for evolutionary changes that would not otherwise occur, or that would otherwise take even longer.